Ozone effects on plants in natural ecosystems.

Tropospheric ozone (O3 ) is an important stressor in natural ecosystems, with well-documented impacts on soils, biota and ecological processes. The effects of O3 on individual plants and processes scale up through the ecosystem through effects on carbon, nutrient and hydrologic dynamics. Ozone effects on individual species and their associated microflora and fauna cascade through the ecosystem to the landscape level. Systematic injury surveys demonstrate that foliar injury occurs on sensitive species throughout the globe. However, deleterious impacts on plant carbon, water and nutrient balance can also occur without visible injury. Because sensitivity to O3 may follow coarse physiognomic plant classes (in general, herbaceous crops are more sensitive than deciduous woody plants, grasses and conifers), the task still remains to use stomatal O3 uptake to assess class and species' sensitivity. Investigations of the radial growth of mature trees, in combination with data from many controlled studies with seedlings, suggest that ambient O3 reduces growth of mature trees in some locations. Models based on tree physiology and forest stand dynamics suggest that modest effects of O3 on growth may accumulate over time, other stresses (prolonged drought, excess nitrogen deposition) may exacerbate the direct effects of O3 on tree growth, and competitive interactions among species may be altered. Ozone exposure over decades may be altering the species composition of forests currently, and as fossil fuel combustion products generate more O3 than deteriorates in the atmosphere, into the future as well.

Disruption of the 'disease triangle' by chemical and physical environmental change.

The physical and chemical environment of the Earth has changed rapidly over the last 100 years and is predicted to continue to change into the foreseeable future. One of the main concerns with potential alterations in climate is the propensity for increases in the magnitude and frequency of extremes to occur. Even though precipitation is predicted to increase in some locations, in others precipitation is expected to decrease and evapotranspiration increase with air temperature, resulting in exacerbated drought in the future. Chemical [ozone (O3 ) and other air contaminants] and subsequent physical alterations in the environment will have a profound effect on the 'disease triangle' (a favourable environment, a susceptible host and a virulent pathogen) and should be included in any analysis of biological response to climate change. The chemical and physical environment affects plant health and alters plant susceptibility to insect and pathogen attack through increased frequency, duration and severity of drought and reduction in host vigour. The potential effects of climate change and O3 on tree diseases with emphasis on the western United States are discussed. We describe a generalised modelling approach to incorporate the complexities of the 'disease triangle' into dynamic vegetation models.

Plasticity in physiological traits in conifers: implications for response to climate change in the western U.S.

Population variation in ecophysiological traits of four co-occurring montane conifers was measured on a large latitudinal gradient to quantitatively assess their potential for response to environmental change. White fir (Abies concolor) had the highest variability, gross photosynthetic rate (Pg), and foliar carbon (C) and nitrogen (N) content. Despite low water use efficiency (WUE), stomatal conductance (gs) of fir was the most responsive to unfavorable environmental conditions. Pinus lambertiana exhibited the least variability in Pg and WUE, and is likely to be the most vulnerable to environmental changes. Pinus ponderosa had an intermediate level of variability, and high needle growth at its higher elevational limits. Pinus Jeffreyi also had intermediate variability, but high needle growth at its southern latitudinal and lower elevational limits. The attributes used to assess tree vigor were effective in predicting population vulnerability to abiotic (drought) and biotic (herbivore) stresses.

Interactive effects of O3 exposure on California black oak (Quercus kelloggii Newb.) seedlings with and without N amendment.

We examined the short-term separate and combined effects of simulated nitrogen (N) deposition (fertilization) and ozone (O(3)) exposure on California black oak seedlings (Quercus kelloggii Newb.), an ecologically important tree of the San Bernardino Mountains downwind of Los Angeles. Realistic concentrations of O(3) were found to cause statistically and biologically significant negative effects on plant health, including lowered photosynthetic ability, lowered water use efficiency, and increased leaf chlorosis and necrosis. When subjected to abrupt changes in light levels, O(3)-exposed plants showed both a slower and smaller response than O(3)-free plants. Fertilized plants exhibited a significantly greater pre- to post-treatment decline in A at saturated [CO(2)] and a significantly lower level of post-treatment chlorosis than unfertilized plants. Fertilization tended to reduce plant sensitivity to O(3).

Isozyme markers associated with O(3) tolerance indicate shift in genetic structure of ponderosa and Jeffrey pine in Sequoia National Park, California.

Effects of canopy ozone (O(3)) exposure and signatures of genetic structure using isozyme markers associated with O(3) tolerance were analyzed in approximately 20-, approximately 80-, and >200-yr-old ponderosa (Pinus ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) in Sequoia National Park, California. For both species, the number of alleles and genotypes per loci was higher in parental trees relative to saplings. In ponderosa pine, the heterozygosity value increased, and the fixation index indicated reduction of homozygosity with increasing tree age class. The opposite tendencies were observed for Jeffrey pine. Utilizing canopy attributes known to be responsive to O(3) exposure, ponderosa pine was more symptomatic than Jeffrey pine, and saplings were more symptomatic than old growth trees. We suggest that these trends are related to differing sensitivity of the two species to O(3) exposure, and to higher O(3) exposures and drought stress that younger trees may have experienced during germination and establishment.

Chronic vs. short-term acute O3 exposure effects on nocturnal transpiration in two Californian oaks.

We tested the effect of daytime chronic moderate ozone (O3) exposure, short-term acute exposure, and both chronic and acute O3 exposure combined on nocturnal transpiration in California black oak and blue oak seedlings. Chronic O3 exposure (70 ppb for 8 h/day) was implemented in open-top chambers for either 1 month (California black oak) or 2 months (blue oak). Acute O3 exposure (approximately 1 h in duration during the day, 120-220 ppb) was implemented in a novel gas exchange system that supplied and maintained known O3 concentrations to a leaf cuvette. When exposed to chronic daytime O3 exposure, both oaks exhibited increased nocturnal transpiration (without concurrent O3 exposure) relative to unexposed control leaves (1.8x and 1.6x, black and blue oak, respectively). Short-term acute and chronic O3 exposure did not further increase nocturnal transpiration in either species. In blue oak previously unexposed to O3, short-term acute O3 exposure significantly enhanced nocturnal transpiration (2.0x) relative to leaves unexposed to O3. California black oak was unresponsive to (only) short-term acute O3 exposure. Daytime chronic and/or acute O3 exposures can increase foliar water loss at night in deciduous oak seedlings.

Stomatal behavior of ozone-sensitive and -insensitive coneflowers (Rudbeckia laciniata var. digitata) in Great Smoky Mountains National Park.

* Morphological and physiological attributes were assessed to elucidate the underlying mechanisms of ozone (O(3)) sensitivity in a highly sensitive species, cutleaf coneflower (Rudbeckia laciniata var. digitata). * Foliage at the same height in the canopy on paired O(3)-sensitive and -insensitive cutleaf coneflowers was assessed for level of foliar symptoms, stomatal density, stomatal responsiveness to dynamic changes in light and leaf-to-air vapor pressure deficit (VPD), steady-state responses to light and CO(2), intrinsic transpirational efficiency, and plant water balance. * There were no morphological differences between the sensitivity types that might have contributed to greater O(3) uptake in sensitive individuals. Stomata of sensitive plants were less responsive than those of insensitive plants to experimentally increased and decreased light intensities, and to increased VPD. O(3)-insensitive plants had greater intrinsic transpirational efficiencies, greater maximum assimilation rates under saturating CO(2) and light, and greater carboxylation rates. * Different physiological attributes vary independently within an individual plant, which collectively confer sensitivity or insensitivity to O(3) injury.

Comparison of calculated and measured foliar O3 flux in crop and forest species.

We designed a new gas exchange system that concurrently measures foliar H2O, O3, and CO2 flux (HOC flux system) while delivering known O3 concentrations. Stomatal responses of three species were tested: snapbean, and seedlings of California black oak (deciduous broadleaf) and blue oak (evergreen broadleaf). Acute O3 exposure (120-250 ppb over an hour) was applied under moderate light and low vapor pressure deficits during near steady state conditions. The rate of stomatal closure was measured when the whole plant was placed in the dark. An adjacent leaf on each plant was also concurrently measured in an O3-free cuvette. Under some conditions, direct measurements and calculated foliar O3 flux were within the same order of magnitude; however, endogenously low gs or O3 exposure-induced depression of gs resulted in an overestimation of calculated O3 fluxes compared with measured O3 fluxes. Sluggish stomata in response to light extinction with concurrent O3 exposure, and incomplete stomatal closure likewise underestimated measured O3 flux.

California black oak response to nitrogen amendment at a high O3, nitrogen-saturated site.

In a nitrogen (N) saturated forest downwind from Los Angeles, California, the cumulative response to long-term background-N and N-amendment on black oak (Quercus kelloggii) was described in a below-average and average precipitation year. Monthly measurements of leaf and branch growth, gas exchange, and canopy health attributes were conducted. The effects of both pollutant exposure and drought stress were complex due to whole tree and leaf level responses, and shade versus full sun leaf responses. N-amended trees had lower late summer carbon (C) gain and greater foliar chlorosis in the drought year. Leaf water use efficiency was lower in N-amended trees in midsummer of the average precipitation year, and there was evidence of poor stomatal control in full sun. In shade, N-amendment enhanced stomatal control. Small differences in instantaneous C uptake in full sun, lower foliar respiration, and greater C gain in low light contributed to the greater aboveground growth observed.

Stomata open at night in pole-sized and mature ponderosa pine: implications for O3 exposure metrics.

Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) is widely distributed in the western USA. We report the lack of stomatal closure at night in early summer for ponderosa pine at two of three sites investigated. Trees at a third site with lower nitrogen dioxide and nitric acid exposure, but greater drought stress, had slightly open stomata at night in early summer but closed stomata at night for the rest of the summer. The three sites had similar background ozone exposure during the summer of measurement (2001). Nighttime stomatal conductance (gs) ranged from one tenth to one fifth that of maximum daytime values. In general, pole-sized trees (< 40 years old) had greater nighttime gs than mature trees (> 250 years old). In late summer, nighttime gs was low (< 3.0 mmol H2O m(-2) s(-1)) for both tree size classes at all sites. Measurable nighttime gs has also been reported in other conifers, but the values we observed were higher. In June, nighttime ozone (O3) uptake accounted for 9, 5 and 3% of the total daily O3 uptake of pole-sized trees from west to east across the San Bernardino Mountains. In late summer, O3 uptake at night was < 2% of diel uptake at all sites. Nocturnal O3 uptake may contribute to greater oxidant injury development, especially in pole-sized trees in early summer.

Variation in morphological and biochemical O3 injury attributes of mature Jeffrey pine within canopies and between microsites.

Crown morphology and leaf tissue chemical and biochemical attributes associated with ozone (O3) injury were assessed in the lower, mid- and upper canopy of Jeffrey pine (Pinus jeffreyi Grev. & Balf.) growing in mesic and xeric microsites in Sequoia National Park, California. Microsites were designated mesic or xeric based on topography and bole growth in response to years of above-average precipitation. In mesic microsites, canopy response to O3 was characterized by thinner branches, earlier needle fall, less chlorotic leaf mottling, and lower foliar antioxidant capacity, especially of the aqueous fraction. In xeric microsites, canopy response to O3 was characterized by higher chlorotic leaf mottling, shorter needles, lower needle chlorophyll concentration, and greater foliar antioxidant capacity. Increased leaf chlorotic mottle in xeric microsites was related to drought stress and increased concurrent internal production of highly reactive oxygen species, and not necessarily to stomatal O3 uptake. Within-canopy position also influenced the expression of O3 injury in Jeffrey pine.

A statistical approach to estimate O3 uptake of ponderosa pine in a mediterranean climate.

In highly polluted sites, stomatal behavior is sluggish with respect to light, vapor pressure deficit, and internal CO2 concentration (Ci) and poorly described by existing models. Statistical models were developed to estimate stomatal conductance (gs) of 40-year-old ponderosa pine at three sites differing in pollutant exposure for the purpose of calculating O3 uptake. Gs was estimated using julian day, hour of day, pre-dawn xylem potential and photosynthetic photon flux density (PPFD). The median difference between estimated and observed field gs did not exceed 10 mmol H2O m(-2) s(-1), and estimated gs within 95% confidence intervals. 03 uptake was calculated from hourly estimated gs, hourly O3 concentration, and a constant to correct for the difference in diffusivity between water vapor and 03. The simulation model TREGRO was also used to calculate the cumulative 03 uptake at all three sites. 03 uptake estimated by the statistical model was higher than that simulated by TREGRO because gas exchange rates were proportionally higher. O3 exposure and uptake were significantly correlated (r2>0.92), because O3 exposure and gs were highly correlated in both statistical and simulation models.

O3 uptake and drought stress effects on carbon acquisition of ponderosa pine in natural stands.

• The effect of O3 exposure or uptake on carbon acquisition (net assimilation (A) or gross photosynthesis (Pg )), with and without drought stress, is reported here in 40-yr-old-ponderosa pine (Pinus ponderosa) trees. • Maximum daily gas exchange was measured monthly for 12 trees at four sites differing in pollutant exposure over two growing seasons with above- and below-average annual precipitation. Gas exchange measures were estimated between sampling periods using a generalized additive regression model. • Both A and Pg generally declined with cumulative O3 exposure or uptake at all sites. As a response variable, Pg was slightly more sensitive than A to cumulative O3 exposure. As a metric, O3 uptake vs exposure permitted slightly better statistical resolution of seasonal response between sites. • The effect of late summer drought stress was statistically significant only at the moderate pollution site, and combined synergistically with O3 exposure or uptake to reduce Pg . The general additive model allows the user to define a deleterious level of cumulative O3 exposure or uptake, and to quantitatively assess biological response.

Seasonal changes in above- and belowground carbohydrate concentrations of ponderosa pine along a pollution gradient.

Seasonal patterns of carbohydrate concentration in coarse and fine roots, stem or bole, and foliage of ponderosa pine (Pinus ponderosa Laws) were described across five tree-age classes from seedlings to mature trees at an atmospherically clean site. Relative to all other tree-age classes, seedlings exhibited greater tissue carbohydrate concentration in stems and foliage, and greater shifts in the time at which maximum and minimum carbohydrate concentration occurred. To determine the effect of environmental stressors on tissue carbohydrate concentration, two tree-age classes (40-year-old and mature) were compared at three sites along a well-established, long-term O3 and N deposition gradient in the San Bernardino Mountains, California. Maximum carbohydrate concentration of 1-year-old needles declined with increasing pollution exposure in both tree-age classes. Maximum fine root monosaccharide concentration was depressed for both 40-year-old and mature trees at the most polluted site. Maximum coarse and fine root starch concentrations were significantly depressed at the most polluted site in mature trees. Maximum bole carbohydrate concentration of 40-year-old trees was greater for the two most polluted sites relative to the cleanest site: the bole appeared to be a storage organ at sites where high O3 and high N deposition decreased root biomass.

Changes in physiological attributes of ponderosa pine from seedling to mature tree.

Plant physiological models are generally parameterized from many different sources of data, including chamber experiments and plantations, from seedlings to mature trees. We obtained a comprehensive data set for a natural stand of ponderosa pine (Pinus ponderosa Laws.) and used these data to parameterize the physiologically based model, TREGRO. Representative trees of each of five tree age classes were selected based on population means of morphological, physiological, and nearest neighbor attributes. Differences in key physiological attributes (gas exchange, needle chemistry, elongation growth, needle retention) among the tree age classes were tested. Whole-tree biomass and allocation were determined for seedlings, saplings, and pole-sized trees. Seasonal maxima and minima of gas exchange were similar across all tree age classes. Seasonal minima and a shift to more efficient water use were reached one month earlier in seedlings than in older trees because of decreased soil water availability in the rooting zone of the seedlings. However, carbon isotopic discrimination of needle cellulose indicated increased water-use efficiency with increasing tree age. Seedlings had the lowest needle and branch elongation biomass growth. The amount of needle elongation growth was highest for mature trees and amount of branch elongation growth was highest for saplings. Seedlings had the highest biomass allocation to roots, saplings had the highest allocation to foliage, and pole-sized trees had the highest allocation to woody tissues. Seedlings differed significantly from pole-sized and older trees in most of the physiological traits tested. Predicted changes in biomass with tree age, simulated with the model TREGRO, closely matched those of trees in a natural stand to 30 years of age.

Use of a single-tree simulation model to predict effects of ozone and drought on growth of a white fir tree.

A physiologically based, single-tree simulation model, TREGRO, was parameterized with existing phenological, allometric, and growth data and used to predict effects of ozone and drought on growth of a 53-year-old white fir (Abies concolor (Gord. & Glend.) Lindl. ex Hildebr.) tree following a 3-year model simulation. Multiple experimental simulations were conducted to assess the individual and interactive effects of ozone (O(3)) exposure and drought on growth of white fir. The effects of O(3) were imposed as reductions in carbon (C) assimilation of 0, 2.5, 5, 10, and 20%. Drought was imposed as 0, 10, 25, and 50% reductions in total annual precipitation. The results of the simulations were compared with the effects of O(3) on white fir seedlings grown in the presence and absence of ozone in open-top chambers and with a field survey of white fir trees subjected to a gradient of O(3). In the O(3) simulations, an O(3)-induced reduction in C assimilation of 2.5% reduced total tree biomass and branch total nonstructural carbohydrate (TNC) content by < 7%. Although quantifiable in simulation experiments, such small reductions would probably not be detectable in the field. Results from both an open-top chamber experiment and a field survey indicated that reductions in C assimilation of white fir growing in elevated O(3) were much greater than 2.5%, but were not statistically different from control values. A simulated O(3) reduction in C assimilation of >/= 10% reduced total tree biomass by 7% and branch TNC by 55%. Results from the field survey indicated that branch elongation was reduced in response to increased O(3) concentration, corroborating the simulated response of reduced C allocation to the branches of white fir. Although simulated reductions in total annual precipitation of >/= 25% reduced final tree biomass, the simulated reductions also reduced O(3) uptake and therefore reduced the O(3) response of white fir. However, a combination of low amounts of O(3) (2.5% reduction in C assimilation) and drought (25% reduction in annual precipitation) synergistically reduced C gain of white fir more than either stress individually. Our simulations predict that moderate drought (no more than a 25% reduction in total annual precipitation) may not ameliorate the response of white fir to O(3) and that moderate amounts of atmospheric O(3) and drought could be more detrimental to white fir than either stress singly.

Response of giant sequoia canopy foliage to elevated concentrations of atmospheric ozone.

We examined the physiological response of foliage in the upper third of the canopy of 125-year-old giant sequoia (Sequoiadendron giganteum Buchholz.) trees to a 61-day exposure to 0.25x, 1x, 2x or 3x ambient ozone concentration. Four branch exposure chambers, one per ozone treatment, were installed on 1-m long secondary branches of each tree at a height of 34 m. No visible symptoms of foliar ozone damage were apparent throughout the 61-day exposure period and none of the ozone treatments affected branch growth. Despite the similarity in ozone concentrations in the branch chambers within a treatment, the trees exhibited different physiological responses to increasing ozone uptake. Differences in diurnal and seasonal patterns of g(s) among the trees led to a 2-fold greater ozone uptake in tree No. 2 compared with trees Nos. 1 and 3. Tree No. 3 had significantly higher CER and g(s) at 0.25x ambient ozone than trees Nos. 1 and 2, and g(s) and CER of tree No. 3 declined with increasing ozone uptake. The y-intercept of the regression for dark respiration versus ozone uptake was significantly lower for tree No. 2 than for trees Nos. 1 and 3. In the 0.25x and 1x ozone treatments, the chlorophyll concentration of current-year foliage of trees Nos. 1 and 2 was significantly higher than that of current-year foliage of tree No. 3. Chlorophyll concentration of current-year foliage on tree No. 1 did not decline with increasing ozone uptake. In all trees, total needle water potential decreased with increasing ozone uptake, but turgor was constant. Although tree No. 2 had the greatest ozone uptake, g(s) was highest and foliar chlorophyll concentration was lowest in tree No. 3 in the 0.25x and 1x ambient atmospheric ozone treatments.

Changes in gas exchange characteristics during the life span of giant sequoia: implications for response to current and future concentrations of atmospheric ozone.

Native stands of giant sequoia (Sequoiadendron giganteum Bucholz) are being exposed to relatively high concentrations of atmospheric ozone produced in urban and agricultural areas upwind. The expected change in environmental conditions over the next 100 years is likely to be unprecedented in the life span (about 2,500 years) of giant sequoia. We determined changes in physiological responses of three age classes of giant sequoia (current-year, 12-, and 125-year-old) to differing concentrations of ozone, and assessed age-related differences in sensitivity to pollutants by examining physiological changes (gas exchange, water use efficiency) across the life span of giant sequoia (current-year, 2-, 5-, 20-, 125-, and > 2,000-year-old trees). The CO(2) exchange rate (CER) was greater in current-year (12.1 micro mol CO(2) m(-2) s(-1)) and 2-year-old seedlings (4.8 micro mol CO(2) m(-2) s(-1)) than in all older trees (3.0 micro mol CO(2) m(-2) s(-1), averaged across the four older age classes). Dark respiration was highest for current-year seedlings (-6.5 +/- 0.7 micro mol CO(2) m(-2) s(-1)) and was increased twofold in symptomatic individuals exposed to elevated ozone concentrations. Stomatal conductance (g(s)) was greater in current-year (355 mmol H(2)O m(-2) s(-1)) and 2-year-old seedlings (200 mmol H(2)O m(-2) s(-1)) than in all older trees (50 mmol H(2)O m(-2) s(-1)), indicating that the ozone concentration in substomatal cavities is higher in young seedlings than in trees. Significant changes in water use efficiency, as indicated by C(i)/C(a), occurred in trees between ages 5 and 20 years. We conclude that giant sequoias seedlings are sensitive to atmospheric ozone until they are about 5 years old. Low conductance, high water use efficiency, and compact mesophyll all contribute to a natural ozone tolerance, or defense, or both, in foliage of older trees.

Physiological adjustment of two full-sib families of ponderosa pine to elevated CO(2).

Seeds from two full-sib families of ponderosa pine (Pinus ponderosa) with known differences in growth rates were germinated and grown in an ambient (350 micro l l(-1)) or elevated (700 micro l l(-1)) CO(2) concentration. Gas exchange at both ambient and elevated CO(2) concentrations was measured 1, 6, 39, and 112 days after the seed coat was shed. Initial stimulation of CO(2) exchange rate (CER) by elevated CO(2) was large (> 100%). On Day 1, CER of seedlings grown in elevated CO(2) and measured at ambient CO(2) was significantly lower than the CER of seedlings grown and measured at ambient CO(2), indicating physiological adjustment of the seedlings exposed to elevated CO(2). Physiological acclimation to elevated CO(2) was complete by Day 39 when there was no significant difference in CER between seedlings grown and measured at ambient CO(2) and seedlings grown and measured at elevated CO(2). After 4 months, the light response of seedlings in the two treatments was determined at both ambient and elevated CO(2). Light compensation point, CER at light saturation, and apparent quantum efficiency of seedlings grown and measured at ambient CO(2) were not significantly different from those of seedlings grown and measured at elevated CO(2). With a short-term increase in CO(2), CER at light saturation (5.16 +/- 0.52 versus 3.13 +/- 0.30 micro mol CO(2) m(-2) s(-1)) and apparent quantum efficiency (0.082 +/- 0.011 versus 0.045 +/- 0.003 micro mol CO(2) micro mol(-1) quanta) were significantly increased. Leaf C/N ratio was significantly increased in the elevated CO(2) treatment. There were few significant differences between families for any response to elevated CO(2). Under the experimental conditions, high growth rate was not correlated with a greater response to elevated CO(2).

Carbon balance in tussock tundra under ambient and elevated atmospheric CO2.

Whole ecosystem CO2 flux under ambient (340 µl/l) and elevated (680 µl/l) CO2 was measured in situ in Eriophorum tussock tundra on the North Slope of Alaska. Elevated CO2 resulted in greater carbon acquisition than control treatments and there was a net loss of CO2 under ambient conditions at this upland tundra site. These measurements indicate a current loss of carbon from upland tundra, possibly the result of recent climatic changes. Elevated CO2 for the duration of one growing season appeared to delay the onset of dormancy and resulted in approximately 10 additional days of positive ecosystem flux. Homeostatic adjustment of ecosystem CO2 flux (sum of species' response) was apparent by the third week of exposure to elevated CO2. Ecosystem dark respiration rates were not significantly higher at elevated CO2 levels. Rapid homeostatic adjustment to elevated CO2 may limit carbon uptake in upland tundra. Abiotic factors were evaluated as predictors of ecosystem CO2 flux. For chambers exposed to ambient and elevated CO2 levels for the duration of the growing season, seasonality (Julian day) was the best predictor of ecosystem CO2 flux at both ambient and elevated CO2 levels. Light (PAR), soil temperature, and air temperature were also predictive of seasonal ecosystem flux, but only at elevated CO2 levels. At any combination of physical conditions, flux of the elevated CO2 treatment was greater than that at ambient. In short-term manipulations of CO2, tundra exposed to elevated CO2 had threefold greater carbon gain, and had one half the ecosystem level, light compensation point when compared to ambient CO2 treatments. Elevated CO2-acclimated tundra had twofold greater carbon gain compared to ambient treatments, but there was no difference in ecosystem level, light compensation point between elevated and ambient CO2 treatments. The predicted future increases in cloudiness could substantially decrease the effect of elevated atmospheric CO2 on net ecosystem carbon budget. These analyses suggest little if any long-term stimulation of ecosystem carbon acquisition by increases in atmospheric CO2.